National Park Service Mojave National Preserve U.S. Department of the Interior

Science Newsletter Blister beetle nest parasites cooperate to mimic the sex pheromone of the solitary bee Habropoda pallida (Hymenoptera: Apidae) Leslie Saul-Gershenz1, Jocelyn G. Millar2, and J. Steven McElfresh2

The solitary bee Habropoda pallida In this Issue: leaves of A. lentiginosus, which leafs out Timberlake is the first native bee to in January (Figure 2). Astragalus emerge in the spring on the Kelso Dunes • Page 1. Blister beetle nest parasites lentiginosus provides nectar to H. pallida, cooperate to mimic the sex pheromone of (Figure 1) in Mojave National Preserve. the solitary bee Habropoda pallida a key pollinator in the Kelso Dunes (Hymenoptera: Apidae) ecosystem and likely also in other dune systems throughout the greater Mojave • Page 5. Functional Ecology Measures: Desert ecosystem. It is considered an Evaluation of Springs in Mojave National Preserve oligolege (specialist) of creosote (Larrea tridentata) (1, 2, 3), but ongoing research • Page 8. Endemic Kelso Dunes Insects: in both low and high precipitation years is a Hotspot of Biodiversity revealing that H. pallida is more plastic in Its emergence is generally synchronized with the onset of blooming of the Borrego milkvetch Astragalus lentiginosus var. borreganus M. E. Jones. This plant is also the sole host plant of adults of the blister beetle Meloe franciscanus (Coleoptera: Meloidae) Van Dyke, an obligate nest parasite of the bee Habropoda pallida. The adult beetles emerge on the dunes in the winter months and feed exclusively on the

Figure 3. Meloe franciscanus larval Figure 1. Kelso Dunes in the Mojave National aggregations averaged 549 larvae (ranged Preserve where Habropoda pallida nests and from 120 to > 2000 and averaged 6.9 +/- 2.8 mm depends on the early blooming nectar source in diameter (range 2-15 mm n=153) 55% were of the Borrego milkvetch, Astragalus ≥7mm) (Saul-Gershenz and Millar 2006). lentiginosus var. borreganus (in the Female abdomens’ average diameter is 6.9 mm foreground), which is also the food plant of the (Hafernik and Saul-Gershenz 2000). Adult adult blister beetle Meloe franciscanus in the female Meloe beetles begin emerging in the winter months. The Mojave Desert ecosystem winter months and lay their eggs at the bases supports 689 species of bees, which is the of grasses. Egg masses laid at the base of highest bee diversity in North America. Image Astragalus sometimes become exposed, due © L. Saul-Gershenz, all rights reserved 2006. to windblown sand movement. Astragalus has 1 University of California, Department of a single tap root system making it vulnerable Figure 2. Adult female Meloe franciscanus to root exposure from wind action while Entomology, One Shields Avenue Davis, CA feeding on Astragalus lentiginosus var. Pleuraphis rigida (big galleta grass) rhizomes 95616, USA. borreganus foliage at Kelso Dunes, Mojave extend radially from the base of the plant 2 National Preserve. Meloe franciscanus is an enabling it to stabilize blowing sand. An University of California, Department of obligate nest parasite of the bee Habropoda oviposition bout ranged from 4.3 to 7.3 hours. Entomology, 3401 Watkins Drive Riverside CA pallida, with beetle larvae completing their Larvae emerged 19 to 43 days later. Published entire development inside a bee’s nest. Image with permission from PNAS 2006, 103:38 92521, USA © L. Saul-Gershenz, all rights reserved 2006. 14039-14044.

Mojave National Preserve Science Newsletter April 2012 This Science Newsletter:

The Mojave Desert is internationally known as a place to conduct scientific research on desert ecosystems. In fact Mojave National Preserve was designated in part to "retain and enhance opportunities for scientific research in undisturbed ecosystems" as stated in the California Desert Protection Act of 1994. Much of this research is conducted through the Sweeney Granite Mountains Desert Research Center, part of the University of California Natural Reserve System, and the Desert Studies Center, operated by the California Desert Studies Consortium of California State Universities. Both are located in the Preserve. Figure 4. The number of male inspections or “visits” representing attraction of male H. pallida bee to hydrocarbon blends mimicking the alkene fractions of female bees, male bees, or triungulins (as per fig. 7) and a control (n=12). Bars marked by the same letter are not The purpose of this newsletter is significantly different (Student-Newman-Keuls procedure, alpha= 0.05). Control (blank) threefold. First, we would like to consisted of hexane. Published with permission from PNAS 2006, 103:38 14039-14044. highlight some of the research being done by scientists in the Preserve and to distribute this information to management and the scientific community. Second, this periodical will allow us to inform the public and research community about science being done by Preserve staff or funded by the National Park Service. And third, we would like to build collaboration between scientists and resource managers so that scientists are made aware of the needs of managers and top quality science is brought to bear on the problems facing resource managers.

This newsletter is published once per year. Copies are available in print at our Kelso Depot Visitor Center, Barstow Headquarters, Desert Studies Center, Sweeney Granite Mountains Desert Research Center, and electronically as pdf documents Figure 5. Habropoda pallida male bee inspecting a M. franciscanus triungulin aggregation on the web1. Articles range from non- emitting a blend of female mimicking sex pheromone. Published with permission from PNAS technical news stories to highly 2006, 103:38 14039-14044. technical research reports. All material in this newsletter has been its host plant preferences than previously of parasitic meloid beetles have different peer-reviewed by subject-matter thought. This relationship is mirrored at strategies for locating the nests of their experts. the Eureka Dunes in Death Valley with A. hosts. Some species climb up on flowers lentiginosus var. micans and at Panamint to await bees when they make a nectar Debra Hughson, Science Advisor Dunes with A. lentiginosus var. variabilis. visit, grabbing onto the bee and being

1 The triungulins (first instar larvae) of the carried back to the nest (4). In some http://www.nps.gov/moja/naturescien ce/sciencenews.htm blister beetle, M. franciscanus are faced meloid species, the beetle larvae crawl with a formidable task: after they emerge, around on the ground in search of bees’ they must find some way of getting to the nests, whereas in others the female nests of their host bees. Different species meloid lays her eggs near the entrance of

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her egg. The nest is 1.5 to 2 m deep in the sand dunes and the next generation of adult blister beetles emerges the following January. The male bees are even attractive to other males when covered with Meloe larvae because they smell like a female bee.

The compositions of the sex pheromone of the female bee and the mimicking M. franciscanus larvae were identified by gas chromatography coupled to mass spectrometry. The sex pheromone is a blend of long-chain cuticular hydrocarbons and is comparatively long lasting (Figure 8) (6). The sex pheromones are produced in the cephalic glands of the female bee. Interestingly, Meloe franciscanus only mimics a subset of the female H. pallida sex pheromone because apparently this subset of compounds is sufficiently attractive to be effective (Figure 4) (6). Thus, these Figure 6. Habropoda pallida male immediately after an encounter with an aggregation of Meloe franciscanus larvae feeding on nectar from Astragalus lentiginosus var. borreganus. beetles have evolved a very clever way of Image © L. Saul-Gershenz, all rights reserved 2004. getting to the scarce resource (a bee’s nest) that they must find in order to a bee’s nest, so the newly hatched larvae field seasons, we determined that the survive in the harsh environment in which just have to crawl into the nest. aggregations of beetle larvae are they live. Instead of searching for a host collectively producing a scent that mimics nest, they attract the host to them, and However, M. franciscanus employs a the volatile sex pheromone utilized by then have the host carry them to a nest. completely different approach. Meloe female bees to attract males for mating. franciscanus eggs from a single large In particular, we found that whole body The Mojave Desert ecosystem is a egg mass all hatch together, emerge extracts from either the female bees or hotspot of native bee diversity with 689 from the sand, and climb up grasses the beetle larvae contained the attractive species, the highest bee diversity in North where there are no flowers with nectar to blend of compounds (Figure 4) (6). America, based on an inventory attract bees (Figure 3). At or close to the conducted in 2004 and 2005 (7). This is top of the stem, the beetle larvae form a Thus, to get to the nest of a host bee, the concordant with the immense botanical tight mass (5). This was the unusual newly hatched M. franciscanus diversity found in the Mojave, as shown behavior that initially caught my (LSG) triungulins aggregate and cooperate to by an inventory of the Mojave National attention. The larvae aggregate and they mimic the sex pheromone of the female Preserve that identified 928 plant species cooperate to stay aggregated for up to bee (Figure 5). When male bees are (8). It is then not surprising to find some fifteen days without feeding (mean 5.4 attracted and approach the aggregations, of the most remarkable examples of days, range 1-15 days, n=36) (5). They the whole mass of several hundred natural selection in this environment move together up and down on big beetle larvae attaches to the male in a where species diversity is high and galetta grass (Pleuraphis rigida) or on fraction of a second (Figure 6) (5, 6). The organisms live at the edge of ecological other species of dune grasses, but male, with all his larvae passengers, then and physiological tolerances and will remain aggregated. Also, their flies off to find a female bee. During likely be under pressure as global climate emergence is timed to coincide with the mating or mating attempts, the beetle shifts. emergence of male H. pallida, which larvae transfer to the female bee, and appear before the females. she carries them back to her nest, where We have determined that M. franciscanus they feed on the pollen and nectar that also parasitizes several other sister In experiments conducted over several she has used to provision the nest, and species of solitary bees in the genera

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2. G.E. Bohart, P.F. Torchio, Y. Maeta, R.W. ndentresearch.htm). Females Rust, Notes on the biology of Emphoropsis 9. E.K. Spangenberg, Plants Eaten By pallida Timberlake. J. Kans. Entomol. Soc. Juvenile Desert Tortoises In The Central 45, 381-92 (1972). Mojave Desert. Paper presented at the 3. P.D. Hurd, E.G. Linsley, The principal National Biological Service Desert Tortoise Larrea bees of the southwestern United Symposium, 1995 (www.tortoise- Triungulins States (Hymenoptera: Apoidea). Smiths tracks.org/publications/ksplants.html). Contrib. Zool. 193, 1-74 (1975). 4. J.D. Pinto, R.B. Selander, The bionomics of Acknowledgements blister beetles of the genus Meloe and a We are deeply indebted to Debra Hughson, Science classification of the New World species. Ill. Advisor for the Mojave National Preserve for her Males Biol. Mon. 42, 1-222 (1970). support and assistance during all aspects of this 5. J. Hafernik, L. Saul-Gershenz, Beetle larvae research as well as Rob Fulton of the Desert Studies cooperate to mimic bees. Nature 405, 35 Center, (CSU). This research is supported by the (2000). Disney Worldwide Conservation Fund, MacBeth 6. L. Saul-Gershenz, J.G. Millar, Phoretic nest Memorial Scholarship grants, and SaveNature.Org. parasites use sexual deceptions to obtain Thanks to Jim Andre and Tasha La Doux for use of transport to their host's nest. Proc. Natl. the facilities at Sweeney Granite Mountain Research Figure 7. Composition of the extracts of Acad. Sci. U.S.A. 103, 14039-44 (2006). Center (UC NRS). A special thanks to Norman female and male H. pallida heads (~n=12) and Meloe franciscanus triungulin aggregations 7. T. Griswold, Personal communication Gershenz for field assistance and support and to (n=6), Numbers equal hydrocarbon chain (2010). John Pinto for his support and continued length. Published with permission from PNAS 8. J.M. André, “Inventory of vascular plants at collaboration and to Rick Westcott. We also thank 2006, 103:38 14039-14044. Mojave National Preserve and Manzanar Peggy Fiedler, John Ascher, Robbin Thorp, Jim Habropoda and Anthophora that are Historic Site” (National Park Service Cane and Neal Williams for their shared expertise found throughout the southern deserts of Inventory and Monitoring Program, 2006, and support during this project. California. The focus of our current www.nps.gov/moja/naturescience/indepe research is the elucidation of the mechanisms of how this sex pheromone mimicry system bridges species and genus boundaries and enables this nest parasite to use multiple bee species as hosts across its extensive range.

In addition, long-term mapping of bee nesting sites in the Mojave National Preserve (LSG) and desert bee biodiversity surveys have yielded two major nesting populations of the bee Calliopsis puellae, which is a specialist pollinator of desert dandelion Malacothrix glabrata in the Preserve, a plant documented as an occasional food plant of juvenile desert tortoises (9). New natural history information has been gathered that will aid in the sustainable management of these pollinators for the future health of the Mojave Desert ecosystem.

References Figure 8. GC-MS analysis and isomeric composition (+/- SD) of the alkene fractions of the 1. E.G. Linsley, The ecology of solitary bees. extracts of the heads of the female and male H. pallida and M. franciscanus larvae revealed that M. franciscanus larvae mimic a subset of the female sex pheromone. Published with Hilgardia 27, 543-99. (1958). permission from PNAS 2006, 103:38 14039-14044.

4 Mojave National Preserve Science Newsletter April 2012 Functional Ecology Measures: Evaluation of Springs in Mojave National Preserve Mark E. Whalon1 and Alexander Johnson1

Within each type of ecosystem there exist limiting factors that regulate maximum primary productivity, species diversity, and control an ecosystem’s food web structure. Water availability is a limiting factor for wetland dependent species in desert ecosystems. Deserts are characterized by the limited amount of rainfall they receive (1), yet numerous small springs and seeps occur in the mountainous desert areas and are focal points of biodiversity. These small wetlands have been historically impacted by cattle grazing and feral burros. Grazing has been retired over much of the Preserve but cattle continue to utilize springs on two active permits in the Preserve, while feral burro management is an ongoing priority. We investigated the use of arthropods (insects) for monitoring selected springs within Mojave National Preserve, hypothesizing that Figure 1. Locations of springs evaluated in this study are shown with respect to Interstate species and numbers present may be 15, Interstate 40, and the boundary of Mojave National Preserve. correlated with cattle or burro use and observable ecosystem impacts (e.g. We sampled arthropods and estimated early April in Mojave National Preserve is water quality degradation). vegetation cover at ten springs (Figure 1, a peak of diurnal activity and seasonal Table 1) in Mojave National Preserve. occurrence of arthropods corresponding Cattle grazing near water sources may Beginning in 2008, we placed ten to at a heat unit accumulation of 250 impact species diversity, disrupt nutrient commercially available yellow sticky traps DD50 °F (degree-days Fahrenheit). We cycling, and alter plant succession at each of the spring locations (Table 1) used various pheromone and kairomone characteristics (2, 3). Our preliminary within or near the watercourse for three odor attractants to lure specific insect work focused on identifying a range of two-day intervals in August and July. species to the traps. The traps were naturally occurring arthropod taxa and Observations were made of vegetation placed as inconspicuously as possible, population density relationships above cover, cattle presence, and existence of taking into consideration terrain, plant and below which detrimental ecological an exclosure fence around the water height, and proximity to well-traveled impacts could be inferred in a spring source. Yellow sticky traps were placed human routes in order to minimize ecosystem by comparing cattle impacted at the same locations in May, 2009, along contact and disturbance by animals and to non-impacted water sources in Mojave with pan traps filled with a water and humans. These techniques of arthropod National Preserve. Terrestrial and aquatic propylene glycol solution, in order to trapping are minimally disruptive and limit arthropods, observations of physical obtain a broader range of specimens in potential confounding factors related to conditions, and water quality parameters better condition for improved human presence. will later be combined in a functional identification. Both pan traps and sticky ecology framework (4, 5) to characterize traps were placed in transects at the four Typical species collected by the sticky the status and level of disturbance of locations with highest biodiversity, based traps and water pan traps included spring ecosystems. on results from previous sampling, during Halictus ligatus (bees), Acanalonia spp. the second week of April, 2010, and (plant hoppers), Ammophila spp. (thread- 1 Department of Entomology, Michigan State collected four days later. The period of waisted wasps), and various members of University, East Lansing, Michigan 48824 USA.

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Chrysididae (cuckoo wasps) (Figure 2). Preliminary identification based on field observations and comparison to known arthropod species from taxonomic records has indicated that in excess of 300 arthropod species were collected near these water sources. A complete list of taxa is pending further identification by specialists in Hymenoptera and dipterans.

Analyses are as yet incomplete and it is unknown what proportion of biodiversity and what species were missed by limited sampling. Our sampling did, however, suggest there may be a higher arthropod diversity at the sites without grazing presence as compared to sites with evidence of cattle impacts, which corresponded to observations of greater vegetation cover, lower water turbidity, and less evidence of trampling at sites without cattle. Existing exclosure fences had been successful at preventing cattle ingress to the water source at the two sampling sites we observed with fences. Figure 2. Examples of insect specimens recovered from the water and sticky traps deployed around Mojave National Preserve springs: Halictus ligatus (top left), plant hopper Greater numbers of pollinator insects and [Acanaloniidae] Acanalonia (top right), [Chrysididae] (bottom left) and Ammophila natural enemy insects (i.e. those which (bottom right). tend to reduce the reproductive success A surprising and significant finding of this western California. After its recent of other insects) were noted in the study was the first detection record in introduction into the U.S., spotted wing undisturbed zones inside the fenced Mojave National Preserve of spotted wing drosophila has been tracked across exclosures. Higher levels of natural drosophila, Drosophila suzukii California, Oregon, Washington, Florida, enemy insects may reduce pestiferous (Matsumura) (Diptera: Drosophilidae), an North Carolina, and Michigan. Six populations such as biting flies and invasive species from Asia (Figure 3) that specimens of spotted wing drosophila mosquitoes in the vicinity of springs. has been destructive in fruit orchards in were collected at the Arrowweed Spring Fewer species overall were sampled at Vegetation Cover (%) water sources utilized by cattle and the Name of water source Date Elev. Excl. Cattle Water Bank Overall spring source areas tended to show more (m) Course signs of cattle trampling. Most of the Arrowweed Spring 1-5 1207 Yes No 90 60 75 water sources utilized by cattle have Bathtub Spring 1-4 1772 No No 20 40 30 pipes and troughs that divert water for Cedar Canyon Rd. Seep 1-4 1558 No Yes 5 livestock. These diversions appear to Ivanpah Spring 1-4 1295 No Yes 15 60 30 reduce flow in the small wetland zones Live Oak Spring 1-4 1512 No No 80 50 70 around spring water sources. Samples Mail Spring 1-5 1527 Yes No 40 60 50 from sites with cattle tended to have Piute Spring 1-4 829 No No 90 45 80 higher levels of biting flies, fecal feeders, Rock House Wash 1-4 1550 No Yes 10 25 15 and gnats compared to sites enclosed by Silver Lead Spring 1-5 1623 No Yes 10 20 15 fences and those without cattle. These Wild Horse Canyon Rd. 1-5 1661 No Yes 30 40 35 initial results, however, represent Table 1. Arthropods were collected at 10 water sources in Mojave National Preserve on sample preliminary indications and are dates: 1 (July 21-22, 2008), 2 (July 28-29, 2008), 3 (Aug. 4-5, 2008), 4 (May 22-23, 2009), and 5 inconclusive pending additional sampling (Apr. 17, 2010). Vegetation cover was estimated within the wet area of the water course, along the riparian bank, and for the spring site overall. Presence of cattle and a fenced exclosure and complete analysis of the data. were noted. 6 Mojave National Preserve Science Newsletter April 2012

sample site (Figure 4) on two of six yellow sticky traps deployed in May, 2010. No spotted wing drosophila were found at Arrowweed spring during the subsequent sampling the following year, but three more specimens were collected in vinegar traps placed at Kelso Depot near a waste container at the visitor parking lot. At the Arrowweed Spring site, where the spotted wing drosophila specimens were initially found, several large trash bags of litter had been left behind by park visitors. Spring flow and vegetation cover at Arrowweed Spring (Figure 4) have increased since retirement of grazing in 1998 and it had one of the higher levels of arthropod biodiversity found during this sampling. Although it is unknown how the spotted wing drosophila arrived in the Preserve, its correlation with trash—especially discarded fruit—was noted. Previously Figure 3- Spotted winged drosophila (SWD) taxonomic characteristics suitable for species infested fruit could account for the identification. Drosophila suzukii, commonly called the spotted-wing drosophila, is a vinegar fly—closely related to Drosophila melanogaster (the common vinegar fly). presence of spotted wing drosophila, although none were found in the garbage left at the site.

Management of high desert springs in Mojave National Preserve to conserve biologic diversity and ecosystem function must take into account potential impacts from visitor use as well as cattle grazing on permitted grazing allotments and feral burros. Periodic inspections of popular locations, such as Arrowweed Spring, for removal of garbage, and monitoring for pestilent insect invasion are recommended. Exclosure fencing appears to function as intended to protect the spring sources area and should continue to be maintained.

References 1. P. Meigs, World distribution of arid and semi-arid homioclimates, in Review of Figure 4. Arrowweed Spring in Mojave National Preserve. Research on Arid Zone Hydrology (Arid Zone Programme, 1, UNESCO, Paris, Using invertebrate bioindicators to assess using invertebrate bioindicators. Aust. J. 1953), pp. 203-210. agricultural sustainability in Australia: Exp. Agr. 47:392-403 (2007). 2. T.L. Fleischner, Ecological costs of livestock proposals and current practices. Aust. J. 5. M.A. McGeoch, The selection, testing, and grazing in Western North America. Conserv. Exp. Agr. 47:379-383 (2007). application of terrestrial insects as Biol. 8, 3:629-444 (1994). 4. K.L. King, K.J. Hutchinson. Pasture and bioindicators. Biol. Rev. 73:181-201 (1998). 3. M.G. Paoletti, L.J. Thomson, A.A. Hoffman, grazing land: assessment of sustainability

7 Mojave National Preserve Science Newsletter April 2012 Endemic Kelso Dunes Insects: a Hotspot of Biodiversity Doug Yanega1 Diversity of several Hymenoptera taxa is concentrated in the North American deserts, with the eastern Mojave Desert their apparent origin and center of diversity, including large bee genera such as Ashmeadiella, Hesperapis, and most notably Perdita (1).

Reference 1. T. Griswold, W. Miller, A Revision of Perdita Polyphylla aeolus, Kelso Dunes endemic Rhaphiomidas tarsalis, Kelso Dunes endemic (Xerophasma) Timberlake (Hymenoptera: (photo by Greg R. Ballmer). (photo by Greg R. Ballmer). Andrenidae). Zootaxa 2517:1-14 (2010). As is true of nearly all of the dune couple of species of weevils including systems in the US, the Kelso Dunes Trigonoscuta kelsoensis, and the aphid This Science Newsletter is a publication of Mojave National Preserve, 2701 Barstow contain a large variety of insects that are wasp Spilomena kelso), suggesting that Road, Barstow, CA 92311. known only from that area. Compiling a we are only just beginning to develop an complete list of endemic insects would be idea of the endemic species that inhabit Editor Debra Hughson, Science Advisor a difficult task given that the various the Kelso Dunes. 760-252-6105, [email protected] species can be in virtually any order, and especially difficult because a fairly large There are some species which are Superintendent Stephanie R. Dubois number have yet to be described or even presently thought not to be endemic, but discovered. The number of practicing which will certainly prove to be so when Contributors insect taxonomists studying the fauna of detailed work is finally complete (e.g., a Leslie Saul-Gershenz the Kelso Dunes is very small (and flightless darkling beetle in the genus Jocelyn G. Millar J. Steven McElfresh always has been), so only the proverbial Lariversius, a newly-described bee in the Mark E. Whalon tip of the iceberg has been scrutinized, genus Hesperapis, and a newly- Alexander Johnson and what *is* known is scattered through described pollen wasp in the genus Doug Yanega the literature, rather than compiled in any Euparagia, all of which are believed to Reviewers single resource. Obviously, the occur elsewhere, though the populations Donald Miller knowledge of each group of insects at Kelso Dunes are morphologically John Pinto Anonymous reflects the taxonomic expertise of those different from those elsewhere, and may few workers who have studied the fauna, only need some genetic work to be Information for Authors and each component has been worked shown as distinct). There are many on in isolation. The crickets, for example, species known to be new, but still The Mojave National Preserve Science Newsletter accepts contributions from received a fair bit of attention many years awaiting formal description, including a qualified researchers on scientific work in ago, resulting in the description of many few species of bees in the genus Perdita, progress or completed in Mojave endemic taxa such as Ammopelmatus and wasps in the genera Belomicrus and National Preserve. Articles can range from general interest stories intended for kelsoensis, Eremopedes kelsoensis, and Plenoculus. Most important, though, is a broad audience to technical research Macrobaenetes kelsoensis. A quick that there are undoubtedly dozens of reports. If you are interested in review also reveals that many of the species still undiscovered, either residing publishing in this Science Newsletter, please contact the editor. Manuscripts, endemic species have only been as specimens in museums, or not yet including figures, photographs, maps, described since 1975, (e.g., the fly having been collected; one can only references, and acknowledgements, Rhaphiomidas tarsalis, the scarabs guess at how many taxa remain to be should be less than 5,000 words. Polyphylla aeolus and Aegialia kelsoi, a found. As such, existing collections of References and notes should be in the Science reference style1. specimens from this area, and future 1 Department of Entomology, Entomology collections, will be of crucial importance if 1http://www.sciencemag.org/about/author Research Museum, University of California, we are ever to even approach some sort s/prep/res/refs.dtl

Riverside, CA 92521-0314 USG of definitive list of endemic species.

8 Mojave National Preserve Science Newsletter April 2012